Spillover from marine reserves related to mechanisms of population regulation

Spillover of fish from marine reserves to adjacent harvested waters may be mediated by density-independent movement, density-dependent movement, or both. If dispersal is by random movement, populations within the reserve must be regulated by density-dependent population growth (DDG). Density-dependent movement (DDM) can also regulate the population if accelerated emigration from a reserve to the surrounding fishing grounds leads to substantially increased mortality. Using spatially explicit models, we show that stock per unit area is bounded for DDG and increases with size for DDM. With DDG, spillover rate per unit area of reserve is maximized with reserves around 50% larger in linear dimension than the minimum size for population persistence. With DDM, spillover per unit area of reserve increases with reserve size. The results highlight the need for the mechanism of population regulation to be incorporated into theoretical and empirical investigations of marine reserve ecology.     

When Is Spillover from Marine Reserves Likely to Benefit Fisheries?

The net movement of individuals from marine reserves (also known as no-take marine protected areas) to the remaining fishing grounds is known as spillover and is frequently used to promote reserves to fishers on the grounds that it will benefit fisheries. Here we consider how mismanaged a fishery must be before spillover from a reserve is able to provide a net benefit for a fishery. For our model fishery, density of the species being harvested becomes higher in the reserve than in the fished area but the reduction in the density and yield of the fished area was such that the net effect of the closure was negative, except when the fishery was mismanaged. The extent to which effort had to exceed traditional management targets before reserves led to a spillover benefit varied with rates of growth and movement of the model species. In general, for well-managed fisheries, the loss of yield from the use of reserves was less for species with greater movement and slower growth. The spillover benefit became more pronounced with increasing mis-management of the stocks remaining available to the fishery. This model-based result is consistent with the literature of field-based research where a spillover benefit from reserves has only been detected when the fishery is highly depleted, often where traditional fisheries management controls are absent. We conclude that reserves in jurisdictions with well-managed fisheries are unlikely to provide a net spillover benefit.     

Large-scale Input Matching by Urban Feral Pigeons (Columba livia)

Ideal Free Distribution (IFD) theory predicts the number of animals choosing habitats of differing quality. Most experimental tests of the IFD have been conducted at small spatial scales (i.e. smaller than maximum daily movement of animals) by comparing the number of animals foraging at adjacent food patches of different quality. Urban pigeons ( Columba livia ) feed in large, open aggregations, and can distribute according to predictions of the IFD at alternative food patches. In this study, we test IFD predictions over a much larger spatial scale by comparing the abundance of urban feral pigeons at four sites spread over the city centre of Montréal, Québec, Canada, to the amount of anthropogenically provided food in each site. We found that the pigeons' distribution among the four sites qualitatively matched that of resources available at these sites. After controlling for the effect of stochastic variation in food resources, two pair-wise comparisons between sites indicated undermatching, one indicated matching and three indicated overmatching of consumers to resources. These results suggest that the pigeons inhabiting the downtown area of Montréal may behave as a single population that distributes qualitatively among foraging sites in proportion to the quantity of food offered, and that deviations from expectations cannot be attributed simply to stochastic variation in the food levels at the sites.     

Environmentally induced dispersal under heterogeneous logistic growth

We consider a single-species model which is composed of several habitats connected by linear migration rates and having logistic growth. A spatially varying, temporally constant environment is introduced by the non-homogeneity of its carrying capacity. Under this condition any type of purely diffusive behavior, characterized in our model by symmetric migration rates, produces an unbalanced population distribution, i.e. some locations receive more individuals than can be supported by the environmental carrying capacity, while others receive less. Using an evolutionarily stable strategy (ESS) approach we show that an asymmetric migration mechanism, induced by the heterogeneous carrying capacity of the environment, will be selected. This strategy balances the inflow and outflow of individuals in each habitat (balanced dispersal), as well as 'balancing' the spatial distribution relative to variation in carrying capacity (the Ideal Free Distribution from habitat selection theory). We show that several quantities are maximized or minimized by the evolutionarily stable dispersal strategy.     

Contribution of triglyceride-rich lipoproteins to plasma free fatty acids.

Free fatty acids are the major lipid fuel of the body. Dysregulation of adipose tissue lipolysis results in increased plasma free fatty acid concentrations, and via that mechanism contributes to insulin resistance in obesity and type 2 diabetes mellitus. Adipose tissue hormone sensitive lipase is thought to be responsible for the production of the majority of free fatty acids. However, a separate contribution comes from the action of endothelial lipases, especially lipoprotein lipase, on triglyceride-rich lipoproteins via a process known as spillover. The primary substrate for spillover appears to be chylomicrons derived from dietary fat. The spillover of fatty acids into the free fatty acid pool varies from one tissue to another. For example, spillover is low ( approximately 14%) in the forearm of healthy volunteers, suggesting that triglyceride fatty acid storage is relatively efficient in skeletal muscle. In contrast, spillover appears to be higher in adipose tissue and may also be higher in the splanchnic bed, based on preliminary data. If systemic spillover is increased in insulin resistant states such as diabetes, this could represent a mechanism contributing to the abnormal increases in plasma concentrations of free fatty acids in that condition.     

Temporal discounting predicts individual competitive success in a human analogue of group foraging

Twenty groups of college undergraduates (N=9-12) participated in a discrete-trials analogue of group foraging in which points, exchangeable for course credit, were the resource to be acquired. Group matching of foragers to patch resource availability was well described by the generalized Ideal Free Distribution, with undermatching (imperfect sensitivity to resource differentials) the norm. Individuals differed on several measures of competitive success, and a measure of temporal discounting (TD) accounted for a significant proportion of the variance in these measures. Tendencies in switch patterns and visit durations differed for the most-impulsive and least-impulsive individuals in a fashion that was consistent with a TD interpretation. When competitive weights, defined in terms of TD scores, were substituted for counts of individuals in the generalized Ideal Free Distribution, group-matching slopes more closely approximated perfect sensitivity than in standard analyses. This was true for groups incorporating all impulsive individuals, all non-impulsive individuals, or a broad range of TD scores. The results suggest that well-understood individual psychological processes are associated with competitive ability in group choice.     

Modelling the Impact of Marine Reserves on a Population with Depensatory Dynamics

In this study, we use a spatially implicit, stage-structured model to evaluate marine reserve effectiveness for a fish population exhibiting depensatory (strong Allee) effects in its dynamics. We examine the stability and sensitivity of the equilibria of the modelled system with regards to key system parameters and find that for a reasonable set of parameters, populations can be protected from a collapse if a small percentage of the total area is set aside in reserves. Furthermore, the overall abundance of the population is predicted to achieve a maximum at a certain ratio \(A\) of reserve area to fished area, which depends heavily on the other system parameters such as the net export rate of fish from the marine reserves to the fished areas. This finding runs contrary to the contested “equivalence at best” result when comparing fishery management through traditional catch or effort control and management through marine reserves. Lastly, we analyse the problem from a bioeconomics perspective by computing the optimal harvesting policy using Pontryagin’s Maximum Principle, which suggests that the value for \(A\) which maximizes the optimal equilibrium fishery yield also maximizes population abundance when the cost per unit harvest is constant, but can increase substantially when the cost per unit harvest increases with the area being harvested.     

Implications of fish home range size and relocation for marine reserve function

Reserves are being used increasingly to conserve fish communities and populations under threat from overfishing, but little consideration has been given to how fish behavior might affect reserve function. This review examines the implications of how fish use space, in particular the occurrence and size of home ranges and the frequency and direction of home range relocations. Examples are drawn primarily from the literature on coral reef fishes, but the principles apply to other habitats. Reserves can protect fish species only if individuals restrict their movements to a localized home range during at least part of the life cycle. Home range sizes increase with body size. In small reserves, a significant proportion of fish whose home ranges are centered within the reserve can be exposed to fishing mortality because their home ranges include non-reserve areas. Relocation of home ranges following initial settlement increases exposure to the fishery, especially if habitat selection is frequency-dependent. Distance, barriers, and costs of movement counter such redistribution. These considerations lead to predictions that population density and mean fish size (1) will form gradients across reserve boundaries with maxima in the center of the reserve and minima outside the reserve away from the boundary; (2) will increase rapidly in newly established reserves, only later providing spillover to adjacent fisheries as density-dependent emigration begins to take effect; and (3) will be higher in reserves that are larger and have higher area:edge ratios, more habitat types, natural barriers between reserve and non-reserve areas, and higher habitat quality inside than outside the reserve. (4) Species with low mobility and weak density-dependence of space use will show the greatest increase in reserves and the strongest benefit for population reproductive capacity, but those with intermediate levels of these traits will provide the greatest spillover benefit to nearby fisheries.     

Vertical distribution of zooplankton: density dependence and evidence for an ideal free distribution with costs

Background  

In lakes with a deep-water algal maximum, herbivorous zooplankton are faced with a trade-off between high temperature but low food availability in the surface layers and low temperature but sufficient food in deep layers. It has been suggested that zooplankton (Daphnia) faced with this trade-off distribute vertically according to an "Ideal Free Distribution (IFD) with Costs". An experiment has been designed to test the density (competition) dependence of the vertical distribution as this is a basic assumption of IFD theory.     

Stock estimation, environmental monitoring and equilibrium control of a fish population with reserve area

For sustainable exploitation of renewable resources, the separation of a reserve area is a natural idea. In particular, in fishery management of such systems, dynamic modelling, monitoring and control has gained major attention in recent years. In this paper, based on the known dynamic model of a fish population with reserve area, the methodology of mathematical systems theory and optimal control is applied. In most cases, the control variable is fishing effort in the unreserved area. Working with illustrative data, first a deterministic stock estimation is proposed using an observer design method. A similar approach is also applied to the estimation of the effect of an unknown environmental change. Then it is shown how the system can be steered to equilibrium in given time, using fishing effort as an open-loop control. Furthermore, a corresponding optimal control problem is also solved, maximizing the harvested biomass while controlling the system into equilibrium. Finally, a closed-loop control model is applied to asymptotically control the system into a desired equilibrium, intervening this time in the reserve area.     

Evolution of heterogeneous perceptual limits and indifference in competitive foraging

The collective behaviour of animal and human groups emerges from the individual decisions and actions of their constituent members. Recent research has revealed many ways in which the behaviour of groups can be influenced by differences amongst their constituent individuals. The existence of individual differences that have implications for collective behaviour raises important questions. How are these differences generated and maintained? Are individual differences driven by exogenous factors, or are they a response to the social dilemmas these groups face? Here I consider the classic case of patch selection by foraging agents under conditions of social competition. I introduce a multilevel model wherein the perceptual sensitivities of agents evolve in response to their foraging success or failure over repeated patch selections. This model reveals a bifurcation in the population, creating a class of agents with no perceptual sensitivity. These agents exploit the social environment to avoid the costs of accurate perception, relying on other agents to make fitness rewards insensitive to the choice of foraging patch. This provides a individual-based evolutionary basis for models incorporating perceptual limits that have been proposed to explain observed deviations from the Ideal Free Distribution (IFD) in empirical studies, while showing that the common assumption in such models that agents share identical sensory limits is likely false. Further analysis of the model shows how agents develop perceptual strategic niches in response to environmental variability. The emergence of agents insensitive to reward differences also has implications for societal resource allocation problems, including the use of financial and prediction markets as mechanisms for aggregating collective wisdom.     

Spillover Effects of a Community-Managed Marine Reserve

The value of no-take marine reserves as fisheries-management tools is controversial, particularly in high-poverty areas where human populations depend heavily on fish as a source of protein. Spillover, the net export of adult fish, is one mechanism by which no-take marine reserves may have a positive influence on adjacent fisheries. Spillover can contribute to poverty alleviation, although its effect is modulated by the number of fishermen and fishing intensity. In this study, we quantify the effects of a community-managed marine reserve in a high poverty area of Northern Mozambique. For this purpose, underwater visual censuses of reef fish were undertaken at three different times: 3 years before (2003), at the time of establishment (2006) and 6 years after the marine reserve establishment (2012). The survey locations were chosen inside, outside and on the border of the marine reserve. Benthic cover composition was quantified at the same sites in 2006 and 2012. After the reserve establishment, fish sizes were also estimated. Regression tree models show that the distance from the border and the time after reserve establishment were the variables with the strongest effect on fish abundance. The extent and direction of the spillover depends on trophic group and fish size. Poisson Generalized Linear Models show that, prior to the reserve establishment, the survey sites did not differ but, after 6 years, the abundance of all fish inside the reserve has increased and caused spillover of herbivorous fish. Spillover was detected 1km beyond the limit of the reserve for small herbivorous fishes. Six years after the establishment of a community-managed reserve, the fish assemblages have changed dramatically inside the reserve, and spillover is benefitting fish assemblages outside the reserve.     

Variation within and among host species in engorgement of larval trombiculid mites

We recovered larval trombiculid mites (i.e. chiggers), vectors of scrub typhus, from small mammal hosts in Taiwan, and compared the relative degree of engorgement (RDE) of the dominant chigger (Leptotrombidium imphalum) from different hosts. Naturally occurring chiggers recovered from Rattus losea and Bandicota indica were 1·4x and 1·3x as engorged as those from Apodemus agrarius. Within each host species, RDE was negatively related to chigger loads, but was mostly unrelated to gender or to body or reproductive condition of hosts. We documented significant variation in chigger engorgement both within and among host species; to the extent that RDE is a proxy for fitness, this contradicts predictions of the Ideal Free Distribution (IFD) that the per capita fitness of vectors should be similar among hosts. Failure to meet predictions of the IFD may reflect the limited mobility of chiggers, which consequently must be less selective in the hosts on which they feed. Further disease control efforts should consider vector feeding success in addition to vector abundance and may be able to capitalize on the unsuitability of certain hosts in supporting disease vectors.     

The ideal free distribution: a review and synthesis of the game-theoretic perspective

The Ideal Free Distribution (IFD), introduced by Fretwell and Lucas in [Fretwell, D.S., Lucas, H.L., 1970. On territorial behavior and other factors influencing habitat distribution in birds. Acta Biotheoretica 19, 16-32] to predict how a single species will distribute itself among several patches, is often cited as an example of an evolutionarily stable strategy (ESS). By defining the strategies and payoffs for habitat selection, this article puts the IFD concept in a more general game-theoretic setting of the “habitat selection game”. Within this game-theoretic framework, the article focuses on recent progress in the following directions: (1) studying evolutionarily stable dispersal rates and corresponding dispersal dynamics; (2) extending the concept when population numbers are not fixed but undergo population dynamics; (3) generalizing the IFD to multiple species.For a single species, the article briefly reviews existing results. It also develops a new perspective for Parker’s matching principle, showing that this can be viewed as the IFD of the habitat selection game that models consumer behavior in several resource patches and analyzing complications involved when the model includes resource dynamics as well. For two species, the article first demonstrates that the connection between IFD and ESS is now more delicate by pointing out pitfalls that arise when applying several existing game-theoretic approaches to these habitat selection games. However, by providing a new detailed analysis of dispersal dynamics for predator-prey or competitive interactions in two habitats, it also pinpoints one approach that shows much promise in this general setting, the so-called “two-species ESS”. The consequences of this concept are shown to be related to recent studies of population dynamics combined with individual dispersal and are explored for more species or more patches.     

The spread of fecally transmitted parasites in socially-structured populations

Mammals are infected by a wide array of gastrointestinal parasites, including parasites that also infect humans and domesticated animals. Many of these parasites are acquired through contact with infectious stages present in soil, feces or vegetation, suggesting that ranging behavior will have a major impact on their spread. We developed an individual-based spatial simulation model to investigate how range use intensity, home range overlap, and defecation rate impact the spread of fecally transmitted parasites in a population composed of social groups (i.e., a socially structured population). We also investigated the effects of epidemiological parameters involving host and parasite mortality rates, transmissibility, disease-related mortality, and group size. The model was spatially explicit and involved the spillover of a gastrointestinal parasite from a reservoir population along the edge of a simulated reserve, which was designed to mimic the introduction pathogens into protected areas. Animals ranged randomly within a "core" area, with biased movement toward the range center when outside the core. We systematically varied model parameters using a Latin hypercube sampling design. Analyses of simulation output revealed a strong positive association between range use intensity and the prevalence of infection. Moreover, the effects of range use intensity were similar in magnitude to effects of group size, mortality rates, and the per-contact probability of transmission. Defecation rate covaried positively with gastrointestinal parasite prevalence. Greater home range overlap had no positive effects on prevalence, with a smaller core resulting in less range overlap yet more intensive use of the home range and higher prevalence. Collectively, our results reveal that parasites with fecal-oral transmission spread effectively in socially structured populations. Future application should focus on parameterizing the model with empirically derived ranging behavior for different species or populations and data on transmission characteristics of different infectious organisms.     

Catastrophes,connectivity and Allee effects in the design of marine reserve networks

Catastrophic events, like oil spills and hurricanes, occur in many marine systems. One potential role of marine reserves is buffering populations against disturbances, including the potential for disturbance-driven population collapses under Allee effects. This buffering capacity depends on reserves in a network providing rescue effects, setting up a tradeoff where reserves need to be connected to facilitate rescue, but also distributed in space to prevent simultaneous extinction. We use a set of population models to examine how dispersal ability and the disturbance regime interact to determine the optimal reserve spacing. We incorporate fishing in a spatially-explicit model to understand the effect of objective choice (e.g. conservation versus fisheries yield) on the optimal reserve spacing. We show that the optimal spacing between reserves increases when accounting for catastrophes with larger spacing needed when Allee effects interact with catastrophes to increase the probability of extinction. We also show that classic tradeoffs between conservation and fishing objectives disappear in the presence of catastrophes. Specifically, we found that at intermediate levels of disturbance, it is optimal to spread out reserves in order to increase both population persistence and to maximize spillover into non-reserve areas.     

Exploitation of a deep-water algal maximum by Daphnia: a stable-isotope tracer study

The exploitation of a deep algal maximum by Daphnia in the absence of fish predation was studied in large indoor mesocosms. Facing the dilemma of low food but high temperature in the epilimnion vs. high food but low temperature in the hypolimnion, Daphnia distribute above and below the thermocline in order to optimise their fitness. Labelling hypolimnetic algae with ¹⁵N revealed that the vertical distribution of Daphnia is dynamic, i.e., all individuals traverse the thermocline and allocate a certain proportion of their time to feeding in the cold water. The overall energy gain from the deep-water algal maximum is lower than from the same algal concentration in the epilimnion due to the low temperature and the limited time an individual spends in the hypolimnion. The results provide mechanistic support for the hypothesis that Daphnia chose their habitat according to an Ideal Free Distribution with Costs model.     

一个设立海洋保护区的捕鱼模型的生物经济分析

考虑一个设立海洋保护区的捕鱼模型．从生物学和经济学两个角度研究海洋保护区的作用．给出所研究系统的生物和经济平衡点以及平衡点的局部稳定性,不稳定性,全局稳定性的条件．研究结果说明设立海洋保护区对于保护海洋资源是一种有效途径,保护区内的鱼群即使在对非保护区进行连续捕捞的情形下也能够保持在一个适当的水平．进一步研究了最优收获策略,并对所得结果的生物经济解释予以说明．     

Marine reserves with ecological uncertainty

To help manage the fluctuations inherent in fish populations scientists have argued for both an ecosystem approach to management and the greater use of marine reserves. Support for reserves includes empirical evidence that they can raise the spawning biomass and mean size of exploited populations, increase the abundance of species and, relative to reference sites, raise population density, biomass, fish size and diversity. By contrast, fishers often oppose the establishment and expansion of marine reserves and claim that reserves provide few, if any, economic payoffs. Using a stochastic optimal control model with two forms of ecological uncertainty we demonstrate that reserves create a resilience effect that allows for the population to recover faster, and can also raise the harvest immediately following a negative shock. The tradeoff of a larger reserve is a reduced harvest in the absence of a negative shock such that a reserve will never encompass the entire population if the goal is to maximize the economic returns from harvesting, and fishing is profitable. Under a wide range of parameter values with ecological uncertainty, and in the ‘worst case’ scenario for a reserve, we show that a marine reserve can increase the economic payoff to fishers even when the harvested population is not initially overexploited, harvesting is economically optimal and the population is persistent. Moreover, we show that the benefits of a reserve cannot be achieved by existing effort or output controls. Our results demonstrate that, in many cases, there is no tradeoff between the economic payoff of fishers and ecological benefits when a reserve is established at equal to, or less than, its optimum size.     

Predicting the population consequences of human disturbance for Ringed Plovers Charadrius hiaticula: a game theory approach

Human disturbance and its potential impacts upon bird populations are currently topical and contentious issues for conservationists. Although many studies have revealed a behavioural impact, or even direct effect on breeding success or survival, these cannot usually be extended to predict the impact on population size. Here we present a population model that allows predictions of the effect that changes in human numbers, visiting a 9-km-long section of the coastline, may have upon the size of a Ringed Plover Charadrius hiaticula population. Human disturbance affects Ringed Plovers in our study area through birds avoiding areas of high disturbance and, in addition, through the accidental trampling of a small number of nests by people walking on the beach. Using the level of human disturbance and habitat variables (which define territory quality) it is possible to predict which areas of beach are occupied and therefore the sites available to the population. Breeding success, for a given area of beach, can be predicted from habitat data. Incorporating known, density-independent, adult mortality allows the equilibrium population size to be predicted. This provides a model that predicts population size. This model is then used to predict the population that the site would support with different, hypothetical, levels of disturbance. If nest loss from human activity was prevented, for example by fencing nests, we predict the Plover population size would increase by 8%. A complete absence of human disturbance would cause a population increase of 85%. If the numbers of people were to double, we predict the population would decrease by 23%.